Some population consequences of variation in preference among individual predators

A number of foraging studies have demonstrated that populations of predators rarely consist of individuals with identical preferences for particular types of prey. Variation among predators can lead to frequency-dependent changes in population preference, because those predators mat prefer the rarer type of prey generally have the greatest influence on population preference. In this study we develop a series of theoretical models which demonstrate how anti-apostatic selection (i.e. selection against the rare form) can arise out of (a) bimodal and (b) normal variation in preference among individuals of the same species. We show that population level anti-apostatic selection can occur even when individual predators show pro-apostatic selection (i.e. selection against the common form). Furthermore, patterns of population prey selection that arise out of variation in preference can potentially be pro-apostatic over one range of relative densities and anti-apostatic over another range of relative densities. Finally, we examine a case study involving predation by female waterboatmen Notonecta glauca and show that the variation in preference in this species is large enough to generate higher anti-apostatic selection than would be expected from the diet selected by the average individual.     

Optimal diet selection,frequency dependence and prey renewal

This paper extends existing models of frequency-dependent diet selection by considering the optimal diet selection of a predator feeding upon prey populations which can be depleted but are also capable of renewal (e.g. immigration, growth, or reproduction). This model and existing models which include prey depletion, predict partial-preference and a generic diet preference for the commonest prey types (apostatic selection). Unlike previous diet selection models, it is found that the optimal diet selection of an individual predator can be to favour the rarest prey type (anti-apostatic selection) when encounter rates are high, even if the individual prey do not differ in their nutritional value. Studies have demonstrated that predators generally show apostatic selection, even when all prey have the same nutritional value. Anti-apostatic selection has also been observed when prey are crowded, and therefore at high density, consistent with the idea of high encounter rates. This anti-apostatic diet selection has previously been proposed as evidence for the use of prey search images by a predator, or variation in individual prey preference. In this paper it is suggested that prey renewal is a further factor, often confounded in experiments, which could favour anti-apostatic selection.     

Differences in food preferences between individuals and populations of domestic cats Felis silvestris catus

The food preferences of 64 cats, of which 28 were domestic pets and 36 were free-ranging animals on three farms, were tested using five food types; a sixth type was also tested on the farm cats. Information was also gathered on the background diet of the house cats (individually) and of each farm cat colony. Consistent differences were found between the predefined groups of cats in their preferences for three foods, Hard Dry (HD), Canned Meat (CM) and Raw Beef (RB), the greatest differences being between house cats and farm cats as a whole. The three colonies of farm cats showed distinct differences in preference, for RB and/or CM, and for the sixth food, Soft Dry (SD). Differences between farms could be explained by a reduced preference for items similar to major components of the background diet of each colony, i.e. selection in favour of foods that were temporally rare. Differences between house and farm cats, and within the house cats, could not be explained in this way; the house cats were neophobic towards RB, and the farm cats ate little of the HD food, possibly because they found it difficult to ingest.     

Using Giving-Up Densities to Test for Dietary Preferences in Primates: An Example with Samango Monkeys ( Cercopithecus (nictitans) mitis erythrarchus)

Teasing apart the components of diet selection is important for understanding an animal’s ecology. We used giving-up densities (GUDs) in artificial food patches to test whether free-ranging samango monkeys (Cercopithecus (nictitans) mitis erythrarchus) treat food and water as complementary resources and to examine dietary preferences. To assess the influence of water on the value of food, we measured harvest of peanuts from food patches augmented with water. To examine dietary preferences, we measured the harvest of peanuts (as a standard for comparing other food classes), raisins, alfalfa pellets, and either mealworms or cat food mixed into sawdust in separate food patches. In addition, we observed the samangos’ order of selection of each food. To differentiate preference from ease of encounter, we measured selectivity for peanuts in triplets of food patches containing 1) peanuts, 2) peanuts mixed with a test food (raisins, alfalfa, or mealworms), and 3) the test food. Water did not influence samango foraging. After peanuts, the samangos treated alfalfa and raisins as approximately equal. The samangos foraged on mealworms lightly and rejected cat food. When each food was mixed with peanuts, the monkeys exhibited an expanding specialist dietary strategy in which they altered their rates of encounter with their preferred foods at high resource densities. Samango monkeys at our study site are not water limited, they consistently favor high-energy foods, and they least often choose animal protein. We conclude that patch-use experiments coupled with direct observations provide a useful means for examining dietary strategy, food preferences, and water limitation.     

Prenatal and early sucking influences on dietary preference in newborn, weaning, and young adult cats

Early experiences are of potential importance in shaping long-term behavior. This study examined the relative influence of prenatal and/or early postnatal experience of chemosensory stimuli on subsequent olfactory and dietary preferences of cats as newborns, at 9-10 weeks, and at 6 months. Cats were exposed to vanillin or 4-ethylguaiacol via their mother's diet either prenatally, postnatally, perinatally (prenatal and postnatal), or experienced no exposure to the stimuli (control). Newborns were given a two-choice olfactory test between the familiar "odor" and no odor; 9-10 week olds were tested for their preference between two food treats, one flavored with the familiar stimulus and the other unflavored; at 6 months, cats were given a choice of two bowls of food, one flavored with the familiar stimulus and the other unflavored. At all ages, cats preferred the familiar, and avoided the unfamiliar, stimulus. Perinatal exposure exerted the strongest influence on preference. Prenatal exposure influenced preference at all ages and postnatal exposure exerted a stronger effect as the cat aged. We conclude that long-term chemosensory and dietary preferences of cats are influenced by prenatal and early (nursing) postnatal experience, supporting a natural and biologically relevant mechanism for the safe transmission of diet from mother to young.     

Nutritional Ecology of <Emphasis Type="Italic">Ateles chamek</Emphasis> in lowland Bolivia: How Macronutrient Balancing Influences Food Choices

All free-living animals must make choices regarding which foods to eat, with the choices influencing their health and fitness. An important goal in nutritional ecology is therefore to understand what governs animals’ diet selection. Despite large variation in the availability of different food items, Peruvian spider monkeys (Ateles chamek) maintain a relatively stable daily protein intake, but allow total energy intake to vary as a function of the composition of available food items. This is referred to as protein-dominated macronutrient balancing. Here we assess the influence of this nutritional strategy on daily and seasonal nutritional intakes, estimate the nutritional value of different foods, and interpret unusual food choices. We conducted continuous all-day observations of focal spider monkeys inhabiting a semideciduous forest in Bolivia. We recorded feeding events, collected foods, and analyzed their nutrient content. By using the Geometric Framework for nutrition, we show that individuals reached their daily end-point in nutrient space —balance between protein and nonprotein energy intake— by consuming nutritionally balanced foods or by alternating between nutritionally complementary foods. The macronutritionally balanced figs of Ficus boliviana were their primary staple food and therefore dominated their overall nutritional intake. Our results also demonstrate that spider monkeys consumed a diverse array of ripe fruits to overcome periods of fig scarcity rather than vice versa; they could obtain sufficient protein on a diet of pure fruit; and unripe figs constituted a nutritionally rewarding and reliable food resource. We hope that the approaches taken and the conclusions reached in this study will catalyze further inquiries into the nutritional ecology of frugivorous primates.     

Comparative analysis of the diet of feral and house cats and wildcat in Europe

Differences in availability of food resources are often manifested in the differentiation of feeding habits of closely related mammal species. Therefore, we assumed that the diet composition and trophic niche of house (i.e., highly dependent on human households), feral (i.e. independent on human households) domestic cats (Felis silvestris catus) and wildcats (Felis s. silvestris) differs. Based on the literature data from Europe (53 study sites), we compared the diet of these three felids analyzed with use of indirect methods (stomach and scat analyses). In the case of the house cat, we additionally compared consumption data obtained directly from prey brought home. Data were expressed as the relative frequency of occurrence to compare dietary patterns. The main prey of the three cat types were small mammals in different ratios. According to the stomach and scat samples, the diet composition of the cat types showed differences in the consumption of rodents, insectivores, wild ungulates, and household food, supporting the “dietary differences originate from varying resources” hypothesis. More opportunistic house cats had a broader trophic niche than feeding specialist wildcats, while feral cats had an intermediate position. The trophic niche breadth of all three cat types increased along a latitudinal gradient from northern to southern areas of Europe. The predation of the house cat which was examined from prey brought home differed from the data obtained by indirect diet analysis; however, it yielded similar results to the diet of the feral cat and the wildcat. Due to their high numbers and similarity of its diet to the wildcat, house cats are a threat to wild animals; therefore, their predation pressure needs to be further investigated.     

Prey selectivity affects reproductive success of a corallivorous reef fish

Most animals consume a narrower range of food resources than is potentially available in the environment, but the underlying basis for these preferences is often poorly understood. Foraging theory predicts that prey selection should represent a trade-off between prey preferences based on nutritional value and prey availability. That is, species should consume preferred prey when available, but select less preferred prey when preferred prey is rare. We employed both field observation and laboratory experiments to examine the relationship between prey selection and preferences in the obligate coral-feeding filefish, Oxymonacanthus longirostris. To determine the drivers of prey selection, we experimentally established prey preferences in choice arenas and tested the consequences of prey preferences for key fitness-related parameters. Field studies showed that individuals fed almost exclusively on live corals from the genus Acropora. While diet was dominated by the most abundant species, Acropora nobilis, fish appeared to preferentially select rarer acroporids, such as A. millepora and A. hyacinthus. Prey choice experiments confirmed strong preferences for these corals, suggesting that field consumption is constrained by availability. In a longer-term feeding experiment, reproductive pairs fed on non-preferred corals exhibited dramatic reductions to body weight, and in hepatic and gonad condition, compared with those fed preferred corals. The majority of pairs fed preferred corals spawned frequently, while no spawning was observed for any pairs fed a non-preferred species of coral. These experiments suggest that fish distinguish between available corals based on their intrinsic value as prey, that reproductive success is dependent on the presence of particular coral species, and that differential loss of preferred corals could have serious consequences for the population success of these dietary specialists.     

The effect of color preferences on the foraging behavior of the green-veined white butterfly (Pieris napi L.)

The color preferences of the green-veined white butterfly Pieris napi (Linnaeus, 1758) that are associated with its foraging behavior were studied in experiments with artificial flower models. It was found that newly emerged inexperienced butterflies that are searching for food prefer flower models that are blue or red and select yellow ones at a considerably lower rate. With repeated visits on artificial flowers, P. napi individuals display are highly consistent in their preference for the first chosen color. Males and females of this species display similar color preferences.     

Nutrient Balancing by Captive Golden Snub-Nosed Monkeys <Emphasis Type="Italic">(Rhinopithecus roxellana</Emphasis>)

An organism’s fitness is tied closely to its ability to obtain food. However, many foods are nutritionally suboptimal on their own, forcing an individual to develop a feeding strategy that actively manages both type and amount of food consumed. Animals in captivity are additionally limited to human provisioned diets, which may be nutritionally inadequate and negatively affect behavior and health. We studied the nutritional intake of captive golden snub-nosed monkeys (Rhinopithecus roxellana) at two locations in China (132 days of continuous, sunrise to sunset focal samples of 7 individuals at each site) and used the Geometric Framework for nutrition to identify their feeding strategy and evaluate diet variation across sites, seasons, and age–sex classes. Captive golden snub-nosed monkeys had a mean nutrient intake of 75% carbohydrates (15% neutral detergent fiber, 60% total nonstructural carbohydrates), 12% fat, and 13% protein (by energy) that differed by location and season owing to differences in the type and amount of food items offered and consumed. Intake at one location differed from that of wild golden snub-nosed monkeys, suggesting that the captive diet was inadequate. These results highlight the importance of developing nutritionally adequate diets for captive animals based on an understanding of their nutritional requirements.     

The plastic fly: the effect of sustained fluctuations in adult food supply on life‐history traits

Many adult traits in Drosophila melanogaster show phenotypic plasticity, and the effects of diet on traits such as lifespan and reproduction are well explored. Although plasticity in response to food is still present in older flies, it is unknown how sustained environmental variation affects life‐history traits. Here, we explore how such life‐long fluctuations of food supply affect weight and survival in groups of flies and affect weight, survival and reproduction in individual flies. In both experiments, we kept adults on constant high or low food and compared these to flies that experienced fluctuations of food either once or twice a week. For these ‘yoyo’ groups, the initial food level and the duration of the dietary variation differed during adulthood, creating four ‘yoyo’ fly groups. In groups of flies, survival and weight were affected by adult food. However, for individuals, survival and reproduction, but not weight, were affected by adult food, indicating that single and group housing of female flies affects life‐history trajectories. Remarkably, both the manner and extent to which life‐history traits varied in relation to food depended on whether flies initially experienced high or low food after eclosion. We therefore conclude that the expression of life‐history traits in adult life is affected not only by adult plasticity, but also by early adult life experiences. This is an important but often overlooked factor in studies of life‐history evolution and may explain variation in life‐history experiments.     

The ‘omnivorous badger dilemma’: towards an integration of nutrition with the dietary niche in wild mammals

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Free-ranging domestic cats (<Emphasis Type="Italic">Felis catus</Emphasis>) on public lands: estimating density,activity, and diet in the Florida Keys

Feral and free-ranging domestic cats (Felis catus) can have strong negative effects on small mammals and birds, particularly in island ecosystems. We deployed camera traps to study free-ranging cats in national wildlife refuges and state parks on Big Pine Key and Key Largo in the Florida Keys, USA, and used spatial capture–recapture models to estimate cat abundance, movement, and activities. We also used stable isotope analyses to examine the diet of cats captured on public lands. Top population models separated cats based on differences in movement and detection with three and two latent groups on Big Pine Key and Key Largo, respectively. We hypothesize that these latent groups represent feral, semi-feral, and indoor/outdoor house cats based on the estimated movement parameters of each group. Estimated cat densities and activity varied between the two islands, with relatively high densities (~4 cats/km²) exhibiting crepuscular diel patterns on Big Pine Key and lower densities (~1 cat/km²) exhibiting nocturnal diel patterns on Key Largo. These differences are most likely related to the higher proportion of house cats on Big Pine relative to Key Largo. Carbon and nitrogen isotope ratios from hair samples of free-ranging cats (n = 43) provided estimates of the proportion of wild and anthropogenic foods in cat diets. At the population level, cats on both islands consumed mostly anthropogenic foods (>80% of the diet), but eight individuals were effective predators of wildlife (>50% of the diet). We provide evidence that cat groups within a population move different distances, exhibit different activity patterns, and that individuals consume wildlife at different rates, which all have implications for managing this invasive predator.     

Best be(e) on low fat: linking nutrient perception,regulation and fitness

Preventing malnutrition through consuming nutritionally appropriate resources represents a challenge for foraging animals. This is due to often high variation in the nutritional quality of available resources. Foragers consequently need to evaluate different food sources. However, even the same food source can provide a plethora of nutritional and non‐nutritional cues, which could serve for quality assessment. We show that bumblebees, Bombus terrestris, overcome this challenge by relying on lipids as nutritional cue when selecting pollen. The bees ‘prioritised’ lipid perception in learning experiments and avoided lipid consumption in feeding experiments, which supported survival and reproduction. In contrast, survival and reproduction were severely reduced by increased lipid contents. Our study highlights the importance of fat regulation for pollen foraging bumblebees. It also reveals that nutrient perception, nutrient regulation and reproductive fitness can be linked, which represents an effective strategy enabling quick foraging decisions that prevent malnutrition and maximise fitness.     

Effects of dietary glucose supplementation on the fasted plasma metabolome in cats and dogs

The study objective was to evaluate nutritional metabolomics in support of companion animal nutrition. The specific purpose was to identify metabolites that differed significantly (q-value < 0.05) in the 23 h fasted plasma metabolome of healthy adult female neutered cats (n = 14) and dogs (n = 14) following 2 weeks adaptation to a single batch of diet, nutritionally complete for both species, with and without dietary glucose supplementation (3.85 % w/w). The study consisted of a two-way, crossover design with samples from individual animals on days 14, 16 and 18 of each diet. Metabolic profiling consistently identified 219 metabolites from cats and 216 metabolites from dogs. Fasted plasma glucose did not change significantly with diet in either species whilst 41 % of cat and 20 % of dog metabolites did change significantly. In general, the two species responded similarly and provided data interpretable within the context of insulin-regulated responses to glucose, for example decreased fatty acid oxidation, increased uptake of branched chain amino acids into muscle, and in cats, decreased glucogenic amino acid catabolism. Other responses, such as increased 1 carbon pool intermediates in cats, were consistent with known nutritional differences between cats and dogs and interpretable within the context of energy metabolism. Data also indicated inter-individual variability and diet-dependent changes in digestion and absorption, providing insights for future study designs. In this proof-of-principle study it was concluded that metabolomics can provide data interpretable to a well-characterised area of nutrition and metabolism and also provide novel insights into metabolic adaptations with relevance to companion animal nutrition.     

Inconsistent use of host plants by the Alaskan swallowtail butterfly: adult preference experiments suggest labile oviposition strategy

Abstract 1. The Alaskan swallowtail butterfly (Papilio machaon aliaska) uses three unrelated plant species as hosts: Cnidium cnidiifolium (Apiaceae), Artemisia arctica (Asteraceae), and Petasites frigidus (Asteraceae). The research presented here investigated whether there are any consistent patterns in host choice by P. m. aliaska females. 2. The first two experiments were designed to test if P. m. aliaska host preference is constant or if it changes from day to day. If host preference is labile, the experiments were designed to also test whether a female’s diet breadth narrows or expands over time. 3. The third experiment tested the host preferences of female offspring from several wild-caught P. m. aliaska females. If P. m. aliaska individuals are specialised in their host use, then all of the offspring from a single female would likely prefer the same host-plant species. This experiment was also designed to test the Hopkins’ host selection principle; does the food plant on which a female is reared as a larva influence her future choices when she is searching for host plants for her own offspring? 4. The results from all of these experiments indicate that P. m. aliaska females vary greatly in their oviposition behaviour and in their preferences for the three host plants. Most populations appear to consist of generalists with labile oviposition behaviour. There is no evidence to support the Hopkins’ host selection principle. 5. It is suggested that the generalised selection of host plants by P. m. aliaska females may be a ‘bet-hedging’ strategy and that this strategy may maximise reproductive fitness in an unpredictable environment.     

Bryophyte use by an insect herbivore: does the crane‐fly Tipula montana select food to maximise growth?

1. Growth rates of immature invertebrates can be of profound importance to subsequent reproductive success in the adult stage. If diet selection in herbivores is a principal means of maximising growth and rate of development, individuals should select food of the highest quality. 2. If disparities are found between performance and preference, these may be the result of individuals balancing nutritional gain against other demands on their behaviour, such as a need for shelter. The responses of the crane‐fly herbivore Tipula montana Curtis (Diptera: Tipulidae) to a range of moss genera from an upland environment were investigated to determine whether diet selection was determined by food quality alone, as measured by its effect on larval growth and survival. 3. Larvae were reared individually on single moss genus diets, and their growth, development, and survival were measured. Each of the mosses supported growth and development although the mean weights achieved differed by a factor of two between food genera. In two‐choice preference experiments, individuals broadly chose bryophyte foods as expected from their quality. One notable exception was the moss Pleurozium schreberi, which gave the best growth performance but was among the least preferred. 4. The two methods of assessing preference gave different results: observations of larvae showed relatively greater selection for dense mosses than faecal pellet analysis. A strong dietary preference for an angiosperm, the sedge Carex bigelowii, over all mosses supported the interpretation that crane‐fly larvae benefit from bryophytes as a refuge, and that this factor may override dietary quality.     

Selection by passerine birds is anti-apostatic at high prey density

Most of the results from past experiments with wild birds and green and brown pastry 'baits' have suggested that disproportionately more of the rare forms are eaten when bait density is high (i.e. selection is anti-apostatic). In two separate series of experiments we presented birds with dishes containing 270 baits of one colour and 30 of another. In series I, five different pairs of colours were presented simultaneously to wild birds at two sites. One colour of each pair was common at one site and the same colour was rare at the other site. After 35 days the ratios of the colours were reversed and the dishes were presented for another 35 days. There was a statistically significant tendency for the colours to be at a higher risk when rare. In series II, three caged blackbirds were offered green and brown baits in two dishes simultaneously; in one dish green was rare and in the other brown was rare. Selection over 6 days was anti-apostatic for all three birds combined but the data proved heterogeneous both between and within individuals. At any one time, each bird tended to concentrate on one colour, irrespective of whether that colour was rare or common. We believe that this behaviour leads to anti-apostatic selection, as has been observed in these and other experiments with pastry prey.     

A Behavioral Mechanism for Incorporating an Unpalatable Food in the Diet of a Generalist Herbivore (Orthoptera: Acrididae)

We investigated mechanisms that could lead to incorporation of unpalatable foods into the diet of a generalist grasshopper, Schistocerca americana: nutritional stress, habituation, learning, and attraction to novelty. The model system involved mesquite, a palatable but inferior food, and mulberry, an unpalatable but adequate food. Nutritional stress, due to prolonged intake of the inferior food, mesquite, did not increase the acceptability of mulberry. Habituation to the deterrent compounds in mulberry and associative learning of the nutritional benefits of mulberry also did not occur. However, mulberry became more acceptable after a day of restriction to a single food type other than mulberry, and even deterrent and nutritionally worthless alternatives such as filter paper became acceptable after a day on any one food type. A tendency to feed on novel food types may be a proximate mechanism for the incorporation of relatively unpalatable, but nutritionally valuable foods into the diet. Novelty and the apparent need for diversity of foods are discussed in the context of exploratory foraging behavior by generalist herbivores.     

Contribution of maternal effects to dietary selection in Mediterranean fruit flies

Individual responses to dietary variation represent a fundamental component of fitness, and nutritional adaptation can occur over just a few generations. Maternal effects can show marked proximate responses to nutrition, but whether they contribute to longer term dietary adaptation is unclear. Here, we tested the hypotheses that maternal effects: (i) contribute to dietary adaptation, (ii) diminish when dietary conditions are constant between generations, (iii) are trait‐specific and (iv) interact with high‐ and low‐quality food. We used experimental evolution regimes in the medfly (Ceratitis capitata) to test these predictions by subjecting an outbred laboratory‐adapted population to replicated experimental evolution on either constant high calorie sugar (‘A’) or low‐calorie starch (‘S’) larval diets, with a standard adult diet across both regimes. We measured the contribution of maternal effects by comparing developmental and adult phenotypes of individuals reared on their own diet with those swapped onto the opposite diet for either one or two generations (high and low maternal effect conditions, respectively), both at the start and after 30 generations of selection. Initially, there were strong maternal effects on female body mass and male mating success but not larval survival. Interestingly, the initial maternal effects observed in female body mass and male mating success showed sex‐specific interactions when individuals from high calorie regimes were tested on low calorie diets. However, as populations responded to selection, the effects of maternal provisioning on all traits diminished. The results broadly supported the predictions. They show how the contribution of maternal effects to dietary responses evolves in a context‐dependent manner, with significant variation across different fitness‐related traits. We conclude that maternal effects can evolve during nutritional adaptation and hence may be an important life history trait to measure, rather than to routinely minimize.